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On the Presence of Late Pleistocene Wapiti

On the Presence of Late Pleistocene Wapiti

On the presence of Late Pleistocene wapiti, canadensis Erxleben, 1777 (Cervidae, Mammalia) in the Palaeolithic site Climăuți II (Moldova) Roman Croitor, Théodor Obada

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Roman Croitor, Théodor Obada. On the presence of Late Pleistocene wapiti, Cervus canadensis Erxleben, 1777 (Cervidae, Mammalia) in the Palaeolithic site Climăuți II (Moldova). Contributions to Zoology, Naturalis, 2018, 87 (1). ￿hal-01737219￿

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HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Contributions to Zoology, 87 (1) 1-10 (2018)

On the presence of Late Pleistocene wapiti, Cervus canadensis Erxleben, 1777 (Cervidae, Mammalia) in the Palaeolithic site Climăuți II (Moldova)

Roman Croitor1, 2, 3, Theodor Obada2

1 Aix-Marseille University, CNRS, UMR 7269, MMSH BP674, 5 Rue Château de l’Horloge, F-13094, Aix-en- Provence, France 2 Institute of Zoology, Academy of Sciences of Moldova, Academiei str. 1, MD-2028, Chisinau, Republic of Moldova 3 E-mail: romancroitor@.com

Keywords: Europe, red , Strongyloceros spelaeus, Cervus elaphus palmidactyloceros, antlers, , systematics.

Abstract elaphus (Flerov, 1952; Sokolov, 1959), but have complicated phylogenetic relationships according to the This article reports antler remains from the Late Paleolithic site modern genetic data (Kuwayama and Ozawa, 2000). of Climăuți II (Republic of Moldova) confirming the presence of wapiti Cervus canadensis in the Late Pleistocene of Western Therefore, our understanding of the “elaphine deer” . The occurrence of wapiti in the East Carpathian area group is more restricted than the definition proposed by 20 ky BP coincides with the local extinction of by Lydekker (1898), who also includes Thorold’s deer giganteus, Crocuta spelaea, and Ursus spelaeus, and substitution of Przewalskum albirostris in his elaphine group. local forest with grazing tundra-steppe Rangifer tarandus Owen (1846) reported and figured an extremely constantini. We here provide an overview of paleontological data and opinions on the presence of Cervus canadensis in Europe, large basal antler portion from the Late Pleistocene a discussion on the taxonomic status and systematic position of deposits of Kent’s Cavern (England) that recalled the extinct deer Cervus elaphus palmidactyloceros, and propose a European morphology but with large dispersal model for wapiti in Europe during the Late Pleistocene. dimensions approaching those of the largest modern North American wapiti antlers. The antler fragment bears two basal tines (brow and bez) as in modern Contents red deer; the circumference of the beam, according to Owen (1846), measures 381 mm (15 inches) between Introduction ...... 1 the brow and the bez tines. The circumference above Material and research methods ...... 4 Systematic Description...... 4 the burr attains 375 mm, thus greatly exceeding the Synonymy...... 4 analogous measurements of modern European red deer Taxonomic remarks...... 4 Cervus elaphus L., which range between 220 and 230 Wapiti remains from Climăuți II...... 6 mm (Friant, 1957). Owen (1846) defined this unusual Discussion...... 7 deer antler from Kent’s Cavern as a representative of Conclusions...... 9 Acknowledgements...... 9 the “round-antlered section of Cervine ” and References...... 9 designated it as a type specimen for a new within the new subgenus Strongyloceros (together with Cervus elaphus, Cervus strongyloceros, and Cervus Introduction canadensis): Cervus (Strongyloceros) spelaeus. This proposed subgenus Strongyloceros thus created the The debates on presence of wapiti-like deer in Europe tautononym Cervus (Strongyloceros) strongyloceros have a long history since Owen (1846) described Schreber, 1836 for the type species of the subgenus. poorly represented fossil remains of an unusually large Owen (1846) assumed a close relationship of this new elaphine deer that rivalled the giant deer Megaloceros species from Kent’s Cavern with North American giganteus in size. Under the term “elaphine deer” we wapiti Cervus strongyloceros Schreber, 1836 and here understand the Eurasian and North American Cervus canadensis Brisson, 1756, with a remark that the deer of the genus Cervus (red deer and wapiti) that deer from Kent’s Cavern may differ in having a longer were traditionally ascribed to the single species Cervus distance between the brow and bez tines. The remains 2 Croitor and Obada – Late Pleistocene wapiti in Moldova

of Sweden. Nonetheless, some authors, for example De Stefano (1911), considered that the presence of Cervus canadensis in Europe cannot be proven based solely on poorly preserved osteological remains. Meanwhile, Cervus elaphus palmidactyloceros De Stefano, 1911 is another large-sized elaphine deer with a pending taxonomical status and systematical position. This intriguing deer from the Late Pleistocene – Early Holocene of Italy is distinguished by its large “wapitoid” body size and the presence of palmated antlers that are unusual for red deer (De Stefano, 1911). Apart from the work of Abbazzi (1995), who regarded C. elaphus palmidactyloceros as an individual morphological variant of European red deer, this poorly understood elaphine deer has enjoyed little scientific attention. Lydekker (1915) regarded Strongyloceros Owen as a junior of Cervus Linnaeus and this viewpoint was generally followed since then. Friant (1952) on the other hand raised Strongyloceros to the generic status and accepted Owen’s species as valid. Apart from the very large size of antlers, Friant (1952, 1957) indicated other diagnostic characteristics of Strongyloceros spelaeus such as the oblique position of antler base with respect to the pedicle, and the complete fusion of the radius and the ulna. It is important to note that Friant Figure 1. The geographical location of the Paleolithic site Climăuți II (indicated with an asterisk). (1957: 61) reported both Sprongyloceros spelaeus and Cervus elaphus in the fauna of the Kent’s Cavern. Friant (1957) also ascribed the remains of a large-sized of Strongyloceros spelaeus and associated fauna come deer from continental Europe, earlier described as from the “terre de la caverne” layer of Würmian Age Cervus primigenius Kaup, to Strongyloceros spelaeus; (Friant, 1957). According to Higham et al. (2011), most even though, as she notes, the antlers of this continental of Kent’s Cavern fauna from the main cave deposits is large-sized stag remained unknown. dated back to 50,000 – 25,000 years before present. According to Lister (1987), some of specimens By the end of 19th century, several reports on the from Kent’s Cavern ascribed by Friant (1952, presence of large fossil wapiti-like deer in Europe 1957) to Strongyloceros spelaeus actually belong appeared in the scientific literature. Gervais (1861, to M. giganteus: the palatal fragment Nr. 5646 with 1872) reported a large-sized Cervus strongyloceros from upper tooth series with weak lingual cingulum in the Cave of Pontil near Saint-Pons (Herault, Southern upper molars (Friant, 1957: Pl. III, fig. 1), the left France) and Loubeau Grotto near Mille (Western hemimandible Nr. 14.11.35 (Friant, 1957: Pl. III, fig. France), associated with a typical Late Pleistocene 3) with well-expressed pachyostosis (the thickness fauna. Belgrand (1883: Pl. 22, fig. 1) described a of mandible in front of the third molar amounts to proximal part of elaphine antler with pedicle from the 37 mm), which falls within the range of variation of Seine valley as “Cervus (canadensis?)” noticing the Megaloceros giganteus samples from Ireland (28.1- exceptionally large size of the antler (the antler beam 41.5 mm) curated at the Natural History Museum of circumference above the bez tine is 210 mm). De Rance London), the metacarpal Nr. 24.1.1938 (Friant, 1957: (1888) reported Strongyloceros spelaeus from Cefn Pl. IV, fig. 6), and the metatarsal Nr. 16.3.1936 (Friant Cave (Wales) in association with a Late Pleistocene 1957: Pl. IV, fig. 7). Indeed, the metapodials from fauna. Geist (1998) indicates the presence of wapiti- Kent’s Cavern perfectly correspond to metapodial like antlers (figured by Ahlén 1965; publication not measurements of the robust short-limbed form of giant available to the authors) in early postglacial deposits deer (Lister, 1994; Croitor et al., 2014). The radius- Contributions to Zoology, 87 (1) – 2018 3

Figure 2. The distal fragment of the left antler of Cervus canadensis from Climăuți II CLM-II-381: A, lateral view; B, upper view; C, view from the broken proximal side. ulna Nr. 5.22, which was regarded by Friant (1957: Pl. the main obstacle for the confirmation of wapiti Cervus( IV, fig. 8) as evidence for the complete fusion of radius canadensis) in Western Eurasia (Friant, 1957). and ulna in the Kent’s Cavern giant stag, actually The present communication proposes a description belongs to a horse (Lister, 1987). Lister (1987), of antler fragments from the Palaeolithic site of however, confirmed that the osteological remains of Climăuți II (Dniester valley, Șoldanești Discrict, the cave stag from Kent’s Cavern are much larger than Republic of Moldova) that undoubtly attest the those of any living European red deer and equal in size presence of wapiti remains in this Late Pleistocene to modern American and Siberian wapiti. Lister (1987) human settlement. also indicated stockier limb bones of the wapiti- The site of Climăuți II (Fig. 1) was discovered in like deer from Kent’s Cavern (“too short for their 1989 and represents a Palaeolithic site, exceptional breadth”: ibidem), which approach the proportions of in Moldova, of specialized mammoth hunters with westernmost living wapiti C. canadensis songaricus remains of a dwelling made of mammoth skulls, tusks, (C. elaphus songaricus according to Lister, 1987). and scapulae (Borziac et al., 2007). The majority of the Finally, the analysis of mitochondrial DNA of fossil remains comes from the upper layer dated elaphine deer remains from Kent’s Cavern revealed that to 20 350 ± 230 y. BP (David and Obada, 1996). The they belong to European red deer Cervus elaphus (Meiri osteological material unearthed from the upper layer of et al., 2013), therefore the question on the presence of the Climăuți site includes 1323 bone fragments, wapiti in the fauna from Kent’s Cavern remains open. the majority of which (83%) belongs to Mammuthus According to Lister (personal communication) the primigenius (19 individuals). The fragments of large presence of wapiti in the fauna of Kent’s Cavern is cervid antlers ascribed to Cervus elaphus by Borziac disproven, since the bones with Cervus elaphus mtDNA et al. (2007) were also retrieved from the upper layer. are large. The diagnostic distal antlers parts are missing These antler fragments bear tool marks and are the only in the European palaeontological record, representing remains representing this cervid at the site, strongly 4 Croitor and Obada – Late Pleistocene wapiti in Moldova

Figure 3. The reconstruction of the wapiti antler CLM-II-381 showing the missing proximal portion and the tines (dashed line): Br, brow (first basal) tine; Bz, bez (second basal) tine; Tz, trez (middle) tine; Cr, crown tine. suggesting that the antler remains have been brought by abbreviations used in the text are: DLM, lateromedial the ancient humans to the site, possibly as raw material diameter; DAP, anteroposterior diameter; L, length, for product manufacture. The exceptionally large size of D, breadth; M, mean value; min, minimum; max, the antler fragments, in comparison to those of modern maximum. red deer, allowed Borziac et al. (2007) to conclude that the elaphine deer from Climăuți II had “des conditions excellentes de vie”. Besides the remarkably large Systematic Description size, the antlers possess some important diagnostic characteristics at the distal portion that permit us to Family Cervidae Goldfuss, 1820 attest to the presence of a new peculiar faunal element Subfamily Goldfuss, 1820 in the Climăuți II site: wapiti (Cervus canadensis). Genus Cervus Linnaeus, 1758 Cervus canadensis Erxleben, 1777

Synonymy Material and research methods 1836: Cervus canadensis Brisson: Schreber, p. 990, tab. 246A. 1836: Cervus strongyloceros Schreber, p. 1074, tabs. 247F-G. 1846: Cervus (Strongyloceros) spelaeus Owen, p. 469, fig. 193. The antler remains from Climăuți II are stored at the 1846: Cervus (Strongyloceros) canadensis Brisson: Owen, Paleontological Museum of the Institute of Zoology of p. 469. the Academy of Sciences of Moldova. The material is 1846: Cervus (Strongyloceros) strongyloceros Schreber: represented by a distal portion of the left antler (CLM- Owen, p. 469. II-381) and a fragment of the left antler from the area of 1853: Cervus intermedius de Serres: Pomel, p. 104. 1883: Cervus (canadensis?) Brisson: Belgrand, p.24, pl. 22, the middle (trez) tine bifurcation (CLM-II-382). Both fig. 1. specimens bear traces of human activity: specimen 1888: Strongyloceros spelaeus Owen: De Rance, p. 11. CLM-II-381 is cut below the middle tine (Fig. 2, C), 1952: Strongyloceros spelaeus Owen: Friant, p. 1398. while specimen CLM-II-382 bears multiple cut marks 2010: Cervus elaphus spelaeus Owen: van der Made, p. 434. and impact pits on the medial side (Fig. 4, C). The systematic description is focused on the morphology of the distal crown, taxonomically Taxonomic remarks meaningful for systematic classification of the genus Cervus at the species and level (Heptner Schreber (1836) granted the authorship of the species and Zalkin, 1947; Flerov, 1952; Geist, 1998). The Cervus canadensis to Brisson (1756). Nonetheless, Contributions to Zoology, 87 (1) – 2018 5

Brisson’s “Regnum Animale” is regarded today as C. elaphus hanglu, C. elaphus affinis, and C elaphus unavailable for zoological taxonomy, since Brisson bactrianus), stressing, however, that the areas of (1756) did not apply binominal zoological nomenclature distribution of Asian wapitis and red deer forms never in his species descriptions. Also, according to the overlap, therefore most probably we are dealing with Article 3.2 of the International Code of Zoological subspecies of the same species. The easy hybridization Nomenclature, no name published before 1758 between wapiti and red deer is also regarded by enters zoological nomenclature. Although a number Heptner and Zalkin (1947) as an argument for including of important generic names are currently accepted various geographical elaphine forms as subspecies in as from Brisson (Gentry, 1994), none of his specific the single species Cervus elaphus. names are accepted as available. Regarding the case of Flerov (1952: 175) included Asian wapitis in the Cervus canadensis in particular, we add that “Cervus subspecies C. elaphus canadensis together with North Canadensis” as mentioned by Brisson (1756) is directly American wapitis. Sokolov (1959) rejected Flerov’s quoted from Ray (1693: p. 84) who used it together with (1952) viewpoint and suggested that the remarkable “Cervus Europaeus” as an indication of the geographic similarity in pelage coloring and antler shape of provenance of the common deer Cervus, not as a Cervus elaphus sibiricus Severtzov and Cervus binominal term. Therefore, these quotations should not elaphus canadensis Erxleben should be regarded as an be regarded as available species designations. evolutionary parallelism and preferred to keep them as Thus, Erxleben (1777) was the first who introduced distinct subspecies of Cervus elaphus Linnaeus. the binominal species name Cervus canadensis for the Schonewald (1994), however, concluded that Canadian stag or wapiti. Schreber (1836) proposed a the differences in behavior (male vocalization), new species name for a large form of North American differences in sexual dimorphism, and semi-lethal deer Cervus strongyloceros accompanied by a brief (less viable) hybridization, support a species rank of description and figures representing a pelage color of Cervus canadensis. a female and a shed antler. In the same work, Schreber Geist (1998) regards wapitis as an eastern radiation (1836) published a description of “Cervus canadensis of red deer that moved into dry, cold, continental Brisson” supplemented with a figure of a stag with regions and adapted to a grazing lifestyle in open fantastic undulated antlers, suggesting that the author landscapes. Although Geist (1998) continues to regard was poorly acquainted with this species. wapitis as a group of subspecies within Cervus elaphus, Severtzoff (1873, 1876) was the first who noticed he confirms an effective reproductive isolation under the striking resemblance between Siberian “Cervus natural conditions between Asian wapitis and the red maral” (not to be confused with Cervus elaphus maral deer subspecies C. elaphus yarkandensis. Although Ogilby currently given to the ) and hybrids between wapiti and European red deer are North American Cervus canadensis, indicating that obtained in captivity, Geist (1998) reports a poor health the only difference between those two cervid forms is condition of such hybrids that likely would cause a low that Canadian deer lack the seasonal change in pelage survival rate under natural conditions. Geist (1998) color (based on an observation of only two captive also reports ethological, physiological, and biological individuals). differences between wapiti and European red deer: the Lydekker (1898) regarded wapiti as a valid species character of rutting call, courtship postures, differences Cervus canadensis and introduced a new subspecies in annual biological cycles, etc. name C. canadensis asiaticus for Asian wapiti which The species rank of Cervus canadensis is supported apparently represents a modification of Cervus maral by recent genetic studies. Kuwayama and Ozawa asiatica as quoted from Severtzoff (1873). According to (2000) revealed a closer phylogenetic relationship Allen (1930), Severtzoff (1873) used the term asiatica of Asian and North American wapiti with in a group sense, not as a technical name, and therefore (Cervus nippon) than with Western (European) red it should be rejected, while the name C. canadensis deer (Cervus elaphus) inferred from mitochondrial asiaticus should then derive from Lydekker (1898). DNA sequences. The mitochondrial DNA analysis Heptner and Zalkin (1947: 69) consider that carried out by Pitra et al. (2004) and Ludt et al. populations of Siberian wapiti in natural conditions (2004) confirmed the monophyletic character of both are genetically isolated from Western and Central Western red deer group and Eastern wapiti group and Asian red deer (i.e. European forms of Cervus elaphus a closer association of wapiti with Cervus nippon and and the Asian subspecies C. elaphus yarkandensis, Przewalskium albirostris. Taking into consideration 6 Croitor and Obada – Late Pleistocene wapiti in Moldova

Table 1. Cervus canadensis from Climăuți II: measurements of morphological pattern typical for Cervus canadensis: the antler CLM-II-381. the antler crown consists of three tines that are situated almost in the same plane (more or less parasagittal Measurements mm with respect to the animal’s main body axis). The first Distance between the trez and first crown tines 410.0 crown tine is the strongest and apparently the longest as well, as is typical for Cervus canadensis. The two DLM of the antler beam between the trez tine 52.1 and the crown distal tines are somewhat inwardly curved (towards the sagittal plane). The antler beam is cylindrical DAP of the antler beam between the trez tine 55.5 and the crown between the middle tine and the distal crown (Table 1); however, the antler becomes somewhat flattened in the D max of the trez tine 43.7 area of bifurcations. The cross-section of the middle D min of the trez tine 42.3 tine is circular, while the crown tines are significantly D max of the beam below the trez tine 74.7 medio-laterally compressed. D min of the beam below the trez tine 55.0 The circumference of the antler beam between the bez and trez tines of specimen CLM-II-381 (Tab. Circumference of the beam below the trez tine 211.0 2) is just slightly below the mean value of antler circumference above the bez tine of the sample of modern C. canadensis from reported by the evidence on reproductive isolation of wapiti and Lydekker (1898) and is slightly larger than the maximal Western red deer populations under natural conditions value of the sample of antlers of modern C. canadensis (Heptner and Zalkin, 1947; Geist, 1998), the from Alaska reported by Boeskorov (2005). It should physiological, ethological, and ecological differences be noted that the sample described by Lydekker (1898) reported by Geist (1998), confirming that Western red includes exceptionally large hunter trophies. The antler deer and wapiti occupy different ecological niches, and from Climăuți II corresponds to the most robust antlers the genetic data opposing Western red deer to wapiti of C. canadensis cherskii from the Late Pleistocene and sika deer, we regard wapiti as a valid species, of Eastern Siberia (Tab. 2). The length of the beam Cervus canadensis, that is clearly distinct from red segment between the trez tine and the first crown deer Cervus elaphus. tine is absolutely and relatively significantly longer in the antler from Climăuți II than in the holotype Wapiti remains from Climăuți II of C. canadensis cherskii (ca. 147 mm; measured from Fig. 2d in Boeskorov, 2005) with respect to the The specimen CLM-II-381 represents a distal portion circumference of the beam segment between the bez of left antler with basal part of middle (trez) tine and and trez tines, which attains 190 mm in the holotype partially preserved crown tines (Figs. 2, 3). The antler of C. canadensis cherskii. The relative length of distal surface bears weak longitudinal ribs and furrows and is beam segment is regarded by Boeskorov (2005) as covered with fine pearling, which is weak compared to one of the diagnostic characters of his subspecies. The that seen in European Cervus elaphus. The middle tine distance between the trez tine and the first crown tine springs off sideward and forms an angle of 80̊ with in the antler from Climăuți is also much longer than the beam. The crown shape represents a comb-like the analogous measurement in C. canadensis fossilis

Table 2. Circumference of the antler beam (mm) between the middle (trez) tine and the crown in fossil and modern Cervus canadensis.

Beam circumference Number of Specimen / sample Geographical provenance Data source (max – M – min) specimens

CLM-II-381 Moldova 176.0 1 This study CLM-II-382 Moldova 206.0 1 This study C. canadensis cherskii Eastern Siberia 165.0 – 186.7 – 210.0 6 Boeskorov (2005) C. canadensis North America 174.6 – 218.1 – 228.6 19 Lydekker (1898) C. canadensis Alaska 168.0 – 187.5 – 210.0 10 Boeskorov (2005) Contributions to Zoology, 87 (1) – 2018 7

Figure 4. The fragment of the left antler of Cervus canaden- sis from Climăuți II CLM- II-382 with basal portion of the broken middle tine: A, lateral side; B, upper side; C, middle side with cut marks.

Zdansky, 1925 (255 mm) from the Pleistocene of advanced and largest C. canadensis canadensis (= C. (Zdansky, 1925: Pl. XV, fig. 1). However, the antler of canadensis sibirica) with large, six-pointed antlers Chinese wapiti has a longer crown part. that occupies mountain biotopes and that is practically The second specimen CLM-II-382 represents a indistinguishable from North American wapiti; the portion of the beam (ca. 37 cm in length) from the area smaller southern form C. canadensis alashanicus of the trez tine bifurcation and belongs to a somewhat with an advanced pattern of rump patch and antlers larger antler (Fig. 4): DLM and DAP of the beam of typical wapiti form but somewhat smaller; and the above the middle tine bifurcation are 64.3 mm and lowland-forest dweller C. canadensis xanthopygus 64.5 mm respectively. The beam circumference above with primitive (or paedomorphic) characters of body the middle tine places this specimen among the most color and social organs, including smaller antlers robust wapiti antlers (Tab. 2). Therefore, the antlers with a short, poorly developed distal portion and a from Climăuți II are as robust and large as the larger poorly distinguished rump patch (Heptner and Zalkin, antlers of modern North American Cervus canadensis 1947; Flerov, 1952; Sokolov, 1959; Geist, 1998). The and exceed the largest specimens of fossil wapiti from northern border of the area of distribution of modern Eastern Siberia in robusticity. Asian wapiti lies in Central Yakutia; however, the palaeontological record indicates a much broader Late Pleistocene area of distribution reaching the far Discussion Northeast (Boeskorov, 2005). Cervus canadensis cherskii from the Late Pleistocene of Eastern Siberia, The modern distribution of Asian C. canadensis ranges is slightly smaller than modern North American C. from Tian-Shan and Altai to the Far East and includes canadensis canadensis, but its antlers are relatively mountainous areas and lowland boreal forests (Flerov, larger (Boeskorov, 2005). Boeskorov (2005) also 1952; Sokolov, 1959). Three subspecies of Asian reports the relatively long distance between bez and wapiti are recognized by Flerov (1952): the most trez tines as a distinguishing taxonomic character of 8 Croitor and Obada – Late Pleistocene wapiti in Moldova

Late Pleistocene Siberian wapiti. Nonetheless, the Lister, 2013). The mitochondrial DNA extracted peculiar antler proportions in Late Pleistocene wapiti from the Romanian sample indicate clearly that the seem to be caused by the shortened distal portion of Romanian elaphine deer falls within the “wapitoid” the antler that approaches C. canadensis cherskii to clade (Parfitt and Lister, 2013; Meiri et al., in press) modern C. canadensis xanthopygus, thus suggesting The confirmed presence of wapiti in Europe is the primitive condition of the short distal part of antlers important for our understanding of the poorly known in these wapitis. The relatively broad metapodials of elaphine deer with palmated antlers from the Late C. canadensis cherskii represent a remarkable affinity Pleistocene and Early Holocene of Italy. Cervus with the robust metapodials of the cave stag from elaphus palmydactyloceros De Stefano, 1911 is a Kent’s Cavern reported by Lister (1987). peculiar and still poorly understood large-sized form The finding of wapiti antlers in the Paleolithic of elaphine deer with distal portions of the antlers site of Climăuți II confirms the dispersal into extending into palmations. The distal antler part Western Eurasia of wapiti, coinciding in Moldova consists of a flattened beam with several tines situated with a local faunal turnover shortly before 20 ky in the parasagittal plane (with the exception of the BP: the local extinction of Megaloceros giganteus, specimen in Tab. XIV, fig. 1 in De Stefano, 1911, where Crocuta spelaea, and Ursus spelaeus, as well as a crown tine is split into a transversally-oriented fork), the substitution of local forest reindeer (Rangifer unlike the multiaxial crown as typical for European tarandus ssp.) with Siberian grazing tundra-steppe red deer. De Stefano (1911) clearly distinguished this subspecies Rangifer tarandus constantini (Croitor, large-sized elaphine deer with palmated antlers from 2018). The poor preservation of the wapiti remains at C. elaphus maral Ogilby, which he also reported from Climăuți II (represented so far only by two humanly- Italy. De Stefano (1911), however, rejected the idea imported specimens) may be explained by the cultural that C. elaphus palmidactyloceros could be related to tradition of mammoth-specialized hunters at Climăuți Cervus canadensis. The elaphine deer remains with (Borziac et al., 2007). Although we have only one palmated antlers come from the Late Pleistocene individual, the relatively longer distal portion of antler and the cold phase of the Younger Dryas in Italy (De from Climăuți II may suggest that we are dealing here Stefano, 1911; Abbazzi, 1995). The body size of De with a quite advanced form of wapiti, more advanced Stefano’s deer is very large: the condylobasal skull than C. canadensis xanthopygus and C. canadensis length, as measured from the photo published by cherskii, however, this conclusion should be taken Abbazzi (1995), is ca. 430 mm and falls within the with caution. The presence of wapiti in South- range of variation of modern Siberian C. canadensis Eastern Europe was earlier attested by the analysis of canadensis (411-444 mm) reported by Flerov (1952). mtDNA sequences from Late Pleistocene remains of C. elaphus palmidactyloceros is characterized by elaphine deer from the Emine-Bair-Khosar Cave in remarkable distal palmations of its antlers with the , Ukraine (Stankovic et al., 2011). According palmation tines situated more or less in the same to these authors, two Crimean specimens dated back plane as in wapiti. Abbazzi (1995) casted doubts to 33,100 ± 400 BP and 42,000 ± 1200 BP belong on the validity of C. elaphus palmidactyloceros, to a wapiti closely related to the modern southern suggesting that the palmated distal portion of antler form C. canadensis songaricus (synonymized with rather represents an individual variation of European C. canadensis canadensis by Flerov, 1952, or with red deer, noticing that some of the specimens ascribed C. canadensis sibirica by Sokolov, 1959). Stankovic by De Stefano (1911) to C. elaphus palmidactyloceros et al. (2011) regard the arrival of wapiti in Crimea are not palmated. Abbazzi (1995) also quotes some as a part of invasion of cold-loving forms into examples of occasional development of palmation Eastern Europe at the end of the Würm II/Würm III under certain conditions in some individuals of wapiti, Interstadial period. According to Stankovic et al. which she regards as a subspecies of Cervus elaphus. (2011), the modern Altai environmental conditions, Indeed, unlike European red deer, wapiti Cervus where Asian wapiti survived, represent the recent canadensis generally grows more flattened antlers that analogue of the environment of the full-glacial period occasionally may develop palmations. The “palmated” of Central Europe. The exceptionally large postcranial variants of antlers are found in some individuals of and dental remains of an elaphine deer (larger than modern Asian wapiti (Heptner and Zalkin, 1946: 25), elaphine remains from Kent’s cavern) were reported as well as in North American wapiti (Nelson, 1902: fig. from the Peştera cu Oase site, Romania (Parfitt and 3). Actually, the variability of palmation in the Italian Contributions to Zoology, 87 (1) – 2018 9 sample does not exclude the validity of the taxon C. References elaphus palmidactyloceros, characterized by a large body size and frequent development of distal antler Abbazzi L. 1995. Occurrence of palmated Cervus elaphus from palmations, while the examples of the occasional Italian Late Pleistocene localities. Atti della Accademia Nazionale dei Lincei, Scienze Fiziche e Naturali, Serie IX, development of palmated antlers in red deer quoted 6(3): 189-206. by Abbazzi (1995) in fact concern Cervus canadensis. Ahlén I. 1965. Studies on the red deer, Cervus elaphus L., in Therefore, we suggest that the large elaphine deer with Scandinavia. 1. History of distribution. Swedish Wildlife, 3: palmated antlers from the Italian Peninsula is a local 1–88. form of wapiti, Cervus canadensis palmidactyloceros Allen GM. 1930. and deer from the Asiatic expeditions. American Museum Novitates, 430: 1-19. De Stefano, 1911. Apparently, C. canadensis Belgrand E. 1883. La Seine : I. Le bassin parisien aux âges palmidactyloceros entered the Italian Peninsula during antéhistoriques. Paris: Imprimerie Nationale. the Last Glacial Maximum and became extinct after Boeskorov GG. 2005. Taxonomic position of the Red Deer the cold phase of Younger Dryas. Cervus elaphus L. (Cervidae, Artiodactyla, Mammalia) from the Neopleistocene of Northeastern Asia. Paleontological Journal, 5 : 73-84. Borziac I, Chirica V, David A. 2007. L’Aurignacien moyen Conclusions et tardif de l’espace carpatique-dnestreen. Le gisement Climauti II. Iași:Institut d’Archeologie, Edition PIM. The distal part of the large elaphine deer antler from Brisson MJ. 1756. Regnum Animale in classes IX distributum, Climăuți II provides important diagnostic characters sive synopsis methodica sistens generalem animalium distributionem in classes IX, & duarum primarum classium, confirming the presence of wapiti Cervus( canadensis) quadrupedum scilicet & cetaceorum, particularem in Western Eurasia during the last glaciation. The divisionem in ordines, sectiones, genera & species. Parisiis: wapiti antler from Climăuți II is characterized by a Bauche. relatively long beam segment between the middle Croitor R. 2018. Paleobiology as a clue to Paleolithic taphonomy: (trez) tine and the first crown tine, regarded here as the case of reindeer hunting in Moldova. Quaternaire 29(1): 81-86. an advanced character opposite to the short distal Croitor R, Stefaniak K, Pawłowska K, Ridush B, Wojtal P, Stach antler part of modern C. canadensis xanthopygus from M. 2014. Giant deer Megaloceros giganteus Blumenbach, the Far East and the Late Pleistocene C. canadensis 1799 (Cervidae, Mammalia) from Palaeolithic of Eastern cherskii from Eastern Siberia. The comb-like crown of Europe. Quaternary International, 326-327: 91-104. the antler from Climăuți II is composed of three tines David A, Obadă Th. 1996. Fauna de mamifere din săpăturile stațiunii paleolitice Climăuți II. Buletinul Academiei de situated in almost the same parasagittal plane. 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